Misleading fruits: The non-monophyly of Pseudopiptadenia and Pityrocarpa supports generic re-circumscriptions and a new genus within mimosoid legumes

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PhytoKeys 205: 239-259 (2022)

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Misleading fruits: The non-monophyly of
Pseudopiptadenia and Pityrocarpa supports generic
re-circumscriptions and a new genus
within mimosoid legumes

Leonardo M. Borges', Peter W. Inglis”, Marcelo F. Simon’,
Pétala Gomes Ribeiro’, Luciano P. de Queiroz?

| Universidade Federal de Séo Carlos, Departamento de Botanica, Rodovia Washington Luis, Km 235, Sto
Carlos, SP 13565-905, Brazil 2. Embrapa Recursos Genéticos e Biotecnologia, Parque Estacdo Biolégica, Caixa
Postal 02372, Brasilia, DE 70770-917, Brazil 3 Universidade Estadual de Feira de Santana, Departamento
de Ciéncias Biolégicas. Av. Transnordestina s.n., Novo Horizonte, Feira de Santana, BA, 44036-900, Brazil

Corresponding author: Leonardo M. Borges (aquitemcaqui@gmail.com)

Academic editor: Colin E. Hughes | Received 15 February 2022 | Accepted 19 May 2022 | Published 22 August 2022

Citation: Borges LM, Inglis PW, Simon ME Ribeiro PG, de Queiroz LP (2022) Misleading fruits: The non-monophyly
of Pseudopiptadenia and Pityrocarpa supports generic re-circumscriptions and a new genus within mimosoid legumes.
In: Hughes CE, de Queiroz LP, Lewis GP (Eds) Advances in Legume Systematics 14. Classification of Caesalpinioideae
Part 1: New generic delimitations. PhytoKeys 205: 239-259. https://doi.org/10.3897/phytokeys.205.82275

Abstract

Generic delimitation in Piptadenia and allies (mimosoid legumes) has been in a state of flux, particularly
caused by over-reliance on fruit and seed morphology to segregate species out of Piptadenia into the genera
Parapiptadenia, Pityrocarpa and Pseudopiptadenia. Although supporting their segregation from Piptadenia,
previous phylogenetic analyses suggested that some of these segregated genera are not monophyletic.
Here, we test the monophyly of Parapiptadenia, Pityrocarpa and Pseudopiptadenia with dense taxon
sampling across these genera, including the type species of each genus. Our analysis recovers Parapitadenia
as monophyletic, but places Pseudopiptadenia species in two distinct lineages, one of which includes all
three species of Pityrocarpa. Given that the type species of both Pseudopiptadenia and Pityrocarpa are
nested in the same clade, we subsume Pseudopiptadenia under the older name Pityrocarpa. The remaining
Pseudopiptadenia species are assigned to the new genus Marlimorimia. Alongside high molecular
phylogenetic support, recognition of Parapiptadenia, Pityrocarpa and Marlimorimia as distinct genera

is also supported by combinations of morphological traits, several of which were previously overlooked.

Keywords

Caesalpinioideae, Fabaceae, Leguminosae, Parapiptadenia, Stryphnodendron clade, tropical America

Copyright Leonardo M. Borges et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

240 Leonardo M. Borges et al. / PhytoKeys 205: 239-259 (2022)

Introduction

Generic delimitation in the mimosoid legumes is being continually revised, notably
across the informal Piptadenia group sensu Lewis and Elias (1981), which included
Anadenanthera Speg., Maicrolobius C. Presl, Mimosa L., Parapiptadenia Brenan,
Piptadenia Benth., Pityrocarpa (Benth.) Britton & Rose, Pseudopiptadenia Rauschert
and Stryphnodendron Mart. Most of the proposed generic re-circumscriptions within
the Piptadenia group have involved segregating species out of Piptadenia, which was
morphologically poorly-defined (Brenan 1955) and is known to be polyphyletic
(Luckow et al. 2003; Jobson and Luckow 2007; Simon et al. 2016; Ribeiro et al. 2018).
While previous phylogenetic and phylogenomic analyses confirm the segregation
of Parapiptadenia, Pityrocarpa and Pseudopiptadenia and place them together with
Stryphnodendron and Microlobius in the Stryphnodendron clade sensu Koenen et al.
(2020), the monophyly of these three genera is still uncertain because of incomplete
taxon sampling in previous analyses (Simon et al. 2016; Koenen et al. 2020; Ringelberg
et al. 2022).

Species of Parapiptadenia, Pityrocarpa and Pseudopiptadenia are trees inhabiting
Neotropical rain forests and seasonally dry tropical forests and woodlands (SDTFWs
sensu Queiroz et al. 2017), with the majority of species in South America and just
two taxa in North America (Pi. obliqua (Pers.) Brenan var. obliqua and Ps. psilostachya
(DC.) G.P. Lewis & M.P. Lima) (Brenan 1955, 1963; Rauschert 1982; Lima and Lima
1984; Lewis and Lima 1991; Queiroz 2009). Their bipinnate leaves vary widely in the
number of pinnae, as well as leaflet number, size and shape. Flowers are pentamerous,
dialipetalous or gamopetalous and arranged in elongated spikes. The diverse fruits and
seeds have been the most prominent traits used to define each genus (Brenan 1955;
Lewis and Elias 1981). Parapiptadenia includes six species with plano-compressed
fruits opening along both sutures (typical legumes) and flat, compressed, narrowly-
winged seeds lacking a pleurogram. Eleven species with similar seeds, but with fol-
licles (fruits splitting along the upper suture only) were placed in Pseudopiptadenia
(Rauschert 1982; Lewis and Lima 1991). The three species in Pityrocarpa, which was
first proposed as a section of Piptadenia (Bentham 1842), differ from the other two
genera by their regularly constricted moniliform legumes and lentiform whitish seeds
with an U-shaped pleurogram (Jobson and Luckow 2007).

The first phylogenetic analysis including these three genera recovered each as
monophyletic, with Pseudopiptadenia contorta (DC.) G.P. Lewis & M.P. Lima and
Ps. psilostachya forming a clade sister to Pityrocarpa (three species sampled), while the
relationship of Parapiptadenia (three species sampled) to other genera was uncertain
(Fig. 1; Jobson and Luckow 2007). The relationships amongst these genera and the
putative monophyly of Pseudopiptadenia were later questioned by analyses with larger
DNA sequence datasets and increased taxon sampling (Simon et al. 2016; Ribeiro et
al. 2018). In these analyses, Parapiptadenia (four species sampled) emerged as sister
to a clade including all sampled species of Pseudopiptadenia (five species, including
Ps. contorta and Ps. psilostachya), except Ps. brenanii G.P. Lewis & M.P. Lima, which

Generic re-circumscriptions and a new genus in the Stryphnodendron clade 241

Microlobius Parapiptadenia Pseudopiptadenia2 Pseudopiptadenia |
Pityrocarpa Stryphnodendron 3
Siryphnodendron 1 Pseudopiptadenia 1 SE oP
‘Pseudopiptadenia 2 Stryphnodendron 1 Parapiptadenia
Parapiptadenia Microlobius Pseudopiptadenia 2
Pityrocarpa
Stryphnodendron 2 Stryphnodendron 2 Pityrocarpa
Microlobius 5 ;
Parapiptadenia Stryphnodenadron 1
Pseudopiptadenia- Stryphnodenaron 2 /Pseudopiptadenia 1 Microlobius
Pityrocarpa Siryphnodendron 1 Stryphnodendron 3 Stryphnodendron 2
Jobson & Luckow 2007 Simon et al. 2016 Ribeiro et al. 2018 Lima et al. 2022

Figure |. Topological differences amongst phylogenetic analyses of the Stryphnodendron clade.

was sister to Pityrocarpa (Fig. 1). This latter clade appeared more closely related to
Stryphnodendron and Microlobius than to the group formed by Parapiptadenia and
Pseudopiptadenia. Phylogenomic analyses with sparse taxonomic sampling recovered
slightly different relationships between these three genera (Fig. 1), but reinforced the
non-monophyly of Pseudopiptadenia (Lima et al. 2022; Ringelberg et al. 2022).

While it is clear that the non-monophyly of Pseudopiptadenia means that taxonomic
adjustments are needed, the type species of the genus, Ps. leptostachya, has not been
included in any previous phylogenetic analyses, raising doubts about its placement
and, hence, about which generic name should be applied to the clade containing that
species. In this study, we infer the phylogenetic relationships between Parapiptadenia,
Pityrocarpa and Pseudopiptadenia using near-complete taxon sampling, including the
type species of all three genera, and re-evaluate the circumscriptions of these genera,
based on the resulting phylogenetic hypothesis.

Materials and methods

Phylogenetic inference

To further test the polyphyly of Pseudopiptadenia indicated by previous studies (Simon
et al. 2016; Ribeiro et al. 2018; Ringelberg et al. 2022) and further investigate sister
group relationships across the Stryphnodendron clade, we carried out phylogenetic
analyses including near-complete sampling of species of Parapiptadenia, Pityrocarpa,
Pseudopiptadenia and allies. Phylogenetic analyses were based on the nuclear ribosomal
5.8S subunit and internal transcribed spacer region (nrITS) and plastid regions matkK
and trnD-trnT. We generated 60 new sequences (21 nrITS, 23 matK, 16 trnD-trnT),
including two accessions of Ps. leptostachya, the type species of Pseudopiptadenia, sam-
pled here for the first time. Published sequences of other members of the Stryphnoden-
dron clade and other genera were obtained from GenBank (Hughes et al. 2003; Simon
et al. 2009; Simon et al. 2016; LPWG 2017; Ribeiro et al. 2018). Sampling com-
prised 60 accessions, including nine species (18 accessions) of Pseudopiptadenia (only

242 Leonardo M. Borges et al. / PhytoKeys 205: 239-259 (2022)

the poorly known Ps. colombiana and Ps. pittieri were not sampled), all three species
of Pityrocarpa (six accessions), all six known species of Parapipitadenia (11 accessions),
plus representatives of the allied genera Microlobius (monospecific; two accessions) and
Stryphnodendron (14 accessions, including members of the three major lineages of this
non-monophyletic genus; see Lima et al. 2022). A selection of mimosoid lineages close-
ly related to the Stryphnodendron clade (Jobson and Luckow 2007; Simon et al. 2016;
Ribeiro et al. 2018; Ringelberg et al. 2022) were included as outgroups. Voucher details
and GenBank accession numbers are provided in Table 1 and in the Suppl. material 1.

Total DNA was extracted from about 20 mg of silica gel-dried leaf material using
a modified CTAB-based protocol (Inglis et al. 2018a). We checked DNA quality and
integrity using agarose gel electrophoresis and DNA quantity and purity estimated by
Nanodrop spectrophotometry (Thermo Scientific). Laboratory procedures, primer se-
quences and amplification protocols followed Inglis et al. (2018b) for nrITS and Simon
et al. (2016) for matK and trnD-trnT. PCR products were prepared for direct Sanger
sequencing using ExoSAP (ThermoFisher) and both DNA strands were sequenced us-
ing the Big Dye v.3.1 kit (Applied Biosystems), using the amplification primers. We
obtained further sequences included in the analysis from GenBank (Table 1).

We assembled contigs using Geneious Prime 2021 (https://www.geneious.com)
and aligned matrices with MAFFT v.7 (Katoh and Standley 2013). Maximum Like-
lihood (ML) phylogenetic analysis was performed using IQ-TREE (Nguyen et al.
2015), using 1000 ultrafast bootstrap replicates to estimate branch support (Hoang
et al. 2018) and models estimated with ModelFinder (Kalyaanamoorthy et al. 2017).
Trees were drawn with FigTree (http://tree.bio.ed.ac.uk/software/figtree/) and rooted
using Lachesiodendron viridiflorum (Kunth) P.G. Ribeiro, L.P. Queiroz & Luckow,
following Ringelberg et al. (2022). Analyses of individual loci produced similar to-
pologies, although the plastid trees were substantially less well-resolved compared to
the nrITS phylogeny. In the absence of major incongruence between individual gene
trees, we inferred phylogenetic relationships with a concatenated dataset (nrITS, matk,
trnD-trnT) containing 3280 bp and 13% of missing data and used it as the basis for
proposing taxonomic rearrangements.

Data Resources

The data underpinning the analysis reported in this paper are deposited in GitHub at
https://doi.org/10.5281/zenodo.6611789

Results and discussion

Our densely sampled phylogenetic analysis recovers Parapitadenia as monophyletic,
reinforces the non-monophyly of Pseudopiptadenia and shows that Pityrocarpa is also
non-monophyletic (Fig. 2). Although the backbone of the phylogeny remains weakly-

243

Generic re-circumscriptions and a new genus in the Stryphnodendron clade

Table |. Voucher information and GenBank accession numbers for taxa used in this study. Newly-

generated sequences are in bold. See the Suppl. material 1 for a digital version.

Taxon Voucher Herbarium nrITS matk trnD-trnT

Inga edulis Matt. Queiroz 13797; HUEFS;K JX870764 AF523078 JQ417383
Pennington 13282
Lachesiodendron viridiflorum (Kunth) PG. Ribeiro, L.P Queiroz & Queiroz 13090 HUEFS MG001274 MG001286 MG001305
Luckow
Microlobius foetidus (Jacq.) M. Sousa & G. Andrade Hughes 2150 FHO KT364047. KT364172 FJ981976
Microlobius foetidus Jacq.) M. Sousa & G. Andrade Macqueen 432 FHO AF458783 AF523095 (No data)
Mimosa palmeri Rose Simon 823 FHO KT364059 KT364212 FJ982142
Mimosa pigra L. Hughes 2414 FHO KT364060 KT364213 FJ982148
Mimosa ursina Matt. Simon 704 CEN KT364061 KT364210 FJ982217
Parapiptadenia blanchetii (Benth.) Vaz & M.P. Lima Queiroz 15358 HUEFS OM575100 ON409904 ON409927
Parapiptadenia blanchetii (Benth.) Vaz & M.P. Lima Thomas 12372 NY OM575099 ON409905 (No data)
Parapiptadenia excelsa (Griseb.) Burkart Hughes 2425 FHO KT364062 KT364160 FJ982235
Parapiptadenia ilheusana G.P. Lewis Neves 1659 RB OM575101 KY046081 ON409928
Parapiptadenia pterosperma (Benth.) Brenan Cardoso 2359 HUEFS OM575102 ON409906 ON409929
Parapiptadenia pterosperma (Benth.) Brenan Ribeiro 902 HUEFS MG001260 ©N409910 MG001292
Parapiptadenia rigida (Benth.) Brenan Marestoni 26 HUEFS MG001261 ON409909 (No data)
Parapiptadenia zehntneri (Harms) M.P. Lima & H.C. Lima Cotarelli 2029 HUEFS OM575104 ON409907 (No data)
Parapiptadenia zehntmeri (Harms) M.P. Lima & H.C. Lima Pereira-Silva 3102 CEN KT364063 KT364063 KT364108
Parapiptadenia zehntmeri (Harms) M.P. Lima & H.C. Lima Queiroz 10974 HUEFS OM575105 ON409908 (No data)
Parapiptadenia zehntmeri (Harms) M.P. Lima & H.C. Lima Queiroz 15692 HUEFS OM575106 KX302341 (No data)
Piptadenia gonoacantha (Matt.) J.E. Macbr. Simon 735 FHO KT364065 DQ790620 FJ982238
Piptadenia stipulacea (Benth.) Ducke Simon 702; Queiroz CEN; KT386296 DQ790634 FJ982239
3115 HUEFS

Pityrocarpa leucoxylon (Barneby & J.W. Grimes) Luckow & R.W. Fernandez 2909 NY (No data) DQ790622 (No data)
Jobson
Pityrocarpa moniliformis (Benth.) Luckow & R.W. Jobson Melo 7518 HUEFS (No data) ON409911 ON409936
Pityrocarpa moniliformis (Benth.) Luckow & R.W. Jobson Queiroz 9084 HUEFS ON191501 ON409912 — (No data)
Pityrocarpa moniliformis (Benth.) Luckow & R.W. Jobson Way 2449 K KT364067 KT364162 FJ982242
Pityrocarpa obliqua (Pers.) Brenan subsp. brasiliensis (G.P. Lewis) Queiroz 12903 HUEFS ON191500 ON409920 — (No data)
Luckow & R.W. Jobson
Pityrocarpa obliqua (Pers.) Brenan subsp. obliqua Macqueen 439 FHO KT364068 KT364206 FJ982243
Pseudopiptadenia bahiana G.P. Lewis & M.P. Lima Melo 138 HUEFS OM575115 ON409916 ON409930
Pseudopiptadenia bahiana G.P. Lewis & M.P. Lima Queiroz 15381 HUEFS MG001277. MG001290 ON409931
Pseudopiptadenia bahiana G.P. Lewis & M.P. Lima Queiroz 15504 HUEFS OM575114 ON409917 ON409932
Pseudopiptadenia brenanii G.P. Lewis & M.P. Lima Borges 680 SPF KT364069 (Nodata) KT364111
Pseudopiptadenia brenanii G.P. Lewis & M.P. Lima Cardoso 2807 HUEFS OM575108 ON409914 ON409937
Pseudopiptadenia brenanii G.P. Lewis & M.P. Lima Harley 56005 HUEFS OM575109 ON409915 (No data)
Pseudopiptadenia brenanii G.P. Lewis & M.P. Lima Queiroz 15585 HUEFS MG001278 ON409913 ON409938
Pseudopiptadenia contorta (DC.) G.P. Lewis & M.P. Lima Queiroz 15507 HUEEFS (No data) KT364155 KT364113
Pseudopiptadenia contorta (DC.) G.P. Lewis & M.P. Lima Queiroz 15582 HUEFS MG001279 KX302348 MG001308
Pseudopiptadenia inaequalis (Benth.) Rauschert Lima 7790 RB OM575111 ON409921 ON409939
Pseudopiptadenia leptostachya (Benth.) Rauschert Lima 8231 RB OM575113 ON409922 ON409940
Pseudopiptadenia leptostachya (Benth.) Rauschert Lima 8326 RB OM575112 ON409923 ON409941
Pseudopiptadenia psilostachya (DC.) G.P. Lewis & M.P. Lima Simon 1245 CEN KT364070 KT364170 KT364114
Pseudopiptadenia schumanniana (Taub.) G.P. Lewis & M.P. Lima Lima 7938 RB OM575110 ON409924 ON409942
Pseudopiptadenia sp. Neves 1675 RB OM575116 ON409918 ON409933
Pseudopiptadenia sp. Ribeiro 351 HUEFS OM575117 ON409919 (No data)
Pseudopiptadenia suaveolens (Miq.) J.W. Grimes = P psilostachya Moacir & Clovis sn IAN OM575119 ON409925 ON409934
Pseudopiptadenia warmingii (Benth.) G.P. Lewis & M.P. Lima Queiroz 12761 HUEFS OM575118 ON409926 ON409935
Senegalia macrostachya (Rchb. ex DC.) Kyal. & Boatwr. Miller 1322 CANB KY688790 KY688920 (No data)
Senegalia nigrescens (Oliv.) PJ.H. Hurter Maurin 255 JRAL JQ265858 GQ872237 (No data)
Stryphnodendron adstringens (Matt.) Coville Souza 29702 ESA KT364072 KT364198 KT364116
Stryphnodendron coriaceum Benth. Scalon 718 ESA (No data) KT364200 KT364120
Stryphnodendron duckeanum Occhioni Simon 1343 CEN KT364076 (Nodata) KT364122
Stryphnodendron fissuratum E.M.O. Martins Lvanauskas sn ESA KT364077. KT364175 KT364124
Stryphnodendron foreroi E.M.O. Martins Assis 1143 SPF KT364079 KT364201 KT364126
Stryphnodendron gracile Rizzini & Heringer Scalon 458 ESA KT364080 KT364177 KT364127
Stryphnodendron obovatum Benth. Scalon 712 ESA KT364081 KT364182 KT364130
Stryphnodendron occhionianum E.M.O. Martins Simon 1597 CEN KT364083. (Nodata) KT364132

244

Taxon
Stryphnodendron paniculatum Poepp.
Stryphnodendron polyphyllum Matt.
Stryphnodendron pulcherrimum (Willd.) Hochr.
Stryphnodendron roseiflorum (Ducke) Ducke
Stryphnodendron rotundifolium Matt.
Stryphnodendron velutinum Scalon

LE 99 Se
0.01 95
ee Mi jalelifaghe.raletievepkTafahetalaisten
91 a
91 OO eres flere ia wtmististert
100 i
100
99 pee enisine
1000] ees
100 Sei
97 eo
95 me ainiel
eh Le,
138 tater Fain
100 ee
80 me
99 Sinise
100 nimin
95 2
100 et Sissies Se
9385 fb iteieeteteiciereraun steyaierete
100 eidinreints
94 Jarertta tyniniaiate
: ae 100 18 aime
: 05 esliaimjaisstisinyuisiaierernis
: 99 ret acmjajerctaieteiers
c itd scam peed gen ae
: 100 Aas
400 Jeena
: 5 i 100 feinfutata (mi eie)mietea mia imiaye ta dimtaeiay sim
: : Ceri it)
: : 1400s A feinyelemetfe.ss
: 99 Pa at

Leonardo M. Borges et al. / PhytoKeys 205: 239-259 (2022)

Voucher Herbarium nrITS matk trnD-trnT
Simon 1058 CEN KT364084 (Nodata) KT364133
Mello-Silva 2659 SPF KT364086 KT364184 KT364136
Simon 980 CEN KT364087. =(Nodata) KT364137
Scalon 728 ESA KT364090 KT364193 KT364143
Scalon 715 ESA KT364094 KT364194 KT364147
Scalon 719 ESA KT364101 KT364187 KT364153
Parapiptadenia zehnineri Cotarelli 2029
Parapiptadenia zehnineri Queiroz 15692
Parapiptadenia zehntneri Queiroz 10974
Parapiptadenia zehnineri Pereira-Silva 3102
Parapiptadenia pterosperma Ribeiro 902
Parapiptadenia pterosperma Cardoso 2359 Pa rapiptadenia

Parapiptadenia blanchetii Queiroz 15358
Parapiptadenia ilheusana
Parapiptadenia blanchetii Thomas 12372
Parapiptadenia rigida

Parapiptadenia excelsa

TYPE

Siryphnodendron adstringens

Siryphnodendron rotundifolium

Siryphnodendron obovatum

Siryphnodendron velutinum

Siryphnodendron gracile

Siryphnodendron polyphyllum

Stryphnodendron roseiflorum

Siryphnodendron foreroi

Stryphnodendron pulcherrimum

Siryphnodendron duckeanum

Siryphnodendron coriaceum

Sitryphnodendron fissuratum

Siryphnodendron occhionianum
Siryphnodendron paniculatum

Microlobius foetidus Hughes 2150

Microlobius foetidus Macqueen 432
Pseudopiptadenia brenanii Cardoso 2807
Pseudopiptadenia brenanii Queiroz 15585
Pseudopiptadenia brenanii Harley 56005
Pseudopiptadenia brenanii Borges 680
Pityrocarpa obliqua Macqueen 439

Pityrocarpa obliqua subsp. brasiliensis Queiroz 13003
Pityrocarpa moniliformis Queiroz 9084
Pityrocarpa moniliformis Way 2449
Pityrocarpa moniliformis Melo 7518
Pityrocarpa leucoxyion
Pseudopiptadenia inaequalis
Pseudopiptadenia schumanniana
Pseudopiptadenia leptostachya Lima 8231
Pseudopiptadenia leptostachya Lima 8326
Mimosa pigra

Mimosa ursina

Mimosa palmeri

Piptadenia gonoacantha

Piptadenia stipulacea

Senegalia macrostachya

Senegalia nigrescens

Inga edulis

Lachesiodendron viridifiorum

Pityrocarpa

TYPE

Figure 2. Phylogeny of the Stryphnodendron clade, based on Maximum Likelihood analysis of the con-

catenated nrITS, matK and trnD-trnT data. Significant ultrafast bootstrap values (> 90%) are given above

branches. The tree was rooted using Lachesiodendron viridiflorum. Scale bar: expected number of changes

per site; dotted branches not to scale.

Generic re-circumscriptions and a new genus in the Stryphnodendron clade 245

supported, the three main clades relevant to the delimitation of genera and the taxo-
nomic decisions proposed here have full (100%) bootstrap support.

The first clade, hereafter referred to as Pseudopiptadenia pro parte, includes
Ps. bahiana G.P. Lewis & M.P. Lima, Ps. contorta, Ps. psilostachya, Ps. warmingii
(Benth.) G.P. Lewis & M.P. Lima and a putative new species yet to be described. The
second clade, hereafter referred to as the Pityrocarpa clade, encompasses the remaining
Pseudopiptadenia species, including the type species of the genus, Ps. leptostachya,
intermixed with accessions of the three species of Pityrocarpa, including Pi. moniliformis
(Benth.) Luckow & R.W. Jobson, the type species of Pityrocarpa.

The placement of Parapiptadenia, Pityrocarpa, and Pseudopiptadenia species in three
distinct lineages and the robustly supported monophyly of Parapiptadenia agree with
previous phylogenetic analyses (Fig. 1; Simon et al. 2016; Ribeiro et al. 2018; Lima et
al. 2022; Ringelberg et al. 2022). However, the relationships amongst these three clades
and other members of the Stryphnodendron clade remain unclear, because of the lack of
support across the backbone of the clade (Figs 1 and 2) and disagreement with previous
analyses. For example, although analyses of nuclear and plastid data (Simon et al. 2016;
Ribeiro et al. 2018) also placed Pseudopiptadenia p.p. and Parapiptadenia in the same
clade, this group could be sister to the remainder of the Stryphnodendron clade (Simon et
al. 2016) or sister to the clade comprising Stryphnodendron and Microlobius (Ribeiro et al.
2018). Phylogenomic analyses based on 997 nuclear genes (Lima et al. 2022; Ringelberg
et al. 2022) placed Pseudopiptadenia p.p. as sister to a group including Stryphnodendron
duckeanum Occhioni f. plus a clade formed by Parapiptadenia and the Pityrocarpa clade.
Furthermore, these nodes across the backbone of the Stryphnodendron clade show high
gene tree conflict (Ringelberg et al. 2022) coinciding with very short branches and weak
support in both conventional and phylogenomic analyses, highlighting the difficulties of
inferring relationships across this part of the mimosoid phylogeny.

Despite uncertainties regarding generic relationships, our results provide an
additional example of how over-reliance on particular traits, in this case fruits and
seeds (Brenan 1955; 1963; Lewis and Elias 1981), may lead to unnatural taxonomies.
Presence of follicles and of flat and winged seeds, which were used to diagnose
Pseudopiptadenia, are respectively shared by most lineages within the Stryphnodendron
clade or homoplastic between Pseudopiptadenia p.p. and members of the Pityrocarpa
clade. All this is not to say that fruits have no taxonomic significance, as the vast
majority of Parapiptadenia species have distinctive legumes with valves plicate above
the seeds, not seen in any other member of the Stryphnodendron clade. Nonetheless,
most species in the Pityrocarpa clade, even though variable in seed morphology (flat
and winged vs. lentiform and wingless), share a number of similarities, including the
position of the extrafloral nectaries between or just below the first pair of pinnae;
few pinnae pairs; inflorescence spikes in general solitary and axillary to coeval
leaves; and bifoliolate seedlings (Fig. 3). These features are not shared with most
Pseudopiptadenia p.p. species, which have extrafloral nectaries on the lower half of
the petiole; many pairs of pinnae; inflorescence spikes arranged in complex efoliate
synflorescences; and pinnate or bipinnate seedlings (see Table 2). Although fairly
homogeneous within the Pityrocarpa clade and Pseudopiptadenia p.p., the characters

246 Leonardo M. Borges et al. / PhytoKeys 205: 239-259 (2022)

Figure 3. Inflorescences of Parapiptadenia, Pityrocarpa and Marlimorimia A Parapiptadenia pterosperma
(Benth.) Brenan showing reddish inflorescences in the axils of coeval leaves B Pa. rigida (Benth.) Brenan
showing yellowish inflorescences and fruits with valves plicate above the seeds C Pityrocarpa brenanii
showing whitish, solitary spikes in the axils of coeval leaves D Pi. moniliformis showing yellowish, solitary
spikes in the axil of a coeval leaf E Marlimorimia bahiana (G.P. Lewis & M.P. Lima) L.P. Queiroz & L.M.
Borges, showing whitish spikes clustered in efoliate terminal pseudoracemes F Marlimorimia sp. showing
yellowish spikes clustered in efoliate terminal pseudoracemes (Photos: A PG Ribeiro; B RT de Queiroz
C-E LP Queiroz; F G Siqueira).

highlighted above sometimes vary amongst and within species, particularly in a
context including Parapiptadenia. For example, solitary inflorescences occur in species

of both Parapiptadenia and the Pityrocarpa clade, while Pseudopiptadenia p.p. species

Generic re-circumscriptions and a new genus in the Stryphnodendron clade 247

Table 2. Morphological comparison amongst Parapiptadenia, Pityrocarpa and Marlimorimia. Traits in
bold highlight diagnostic features separating Pityrocarpa and Marlimorimia. EFN - Extrafloral nectary.

Parapiptadenia Pityrocarpa Marlimorimia
Pinnae number 1-8 1-4 (rarely to 5 in Pi. brenanii) 5-10 or more (2-5 in M. bahiana and
M. colombiana)
Petiolar EFN Variable across species Between or just below the first | Between the base and the middle of the petiole
position pair of pinnae
Spike arrangement | Solitary, axillary or supra-axillary | Solitary (very rarely up to 2 in 2-many fasciculate and further arranged in
to coeval leaves Pi. moniliformis), axillary to | efoliate terminal pseudoracemes or on efoliate
coeval leaves nodes below mature leaves
Petals Reddish (yellowish in Pa. excelsa | White to yellowish or greenish; White to yellowish or greenish; united and
and Pa. rigida); united at the base] free and glabrous (united in pubescent
(free in Pa. rigida) Pi. leucoxylon)
Fruit type Legume Follicle Follicle
(dehiscence)
Fruit shape Flat compressed, valves plicate Flat compressed, valves not Flat compressed, valves not plicate above the seeds
above the seeds (except in Pa. plicate above the seeds
excelsa)

Fruit margins Straight to shallowly sinuous Deeply constricted Straight, (shallowly and irregularly sinuous in

(moniliform) (sinuous in M. bahiana, M. colombiana and M. warmingii),

P. brenanii). sometimes constricted where seeds abort

Valve consistency ‘Thin, chartaceous, glabrous Thick, coriaceous, mostly Thin, coriaceous (thicker and harder in
and indumentum pubescent M. warmingii), glabrous
Embryo plumule Developed and multifid Rudimentary (developed and Developed and multifid

multifid in R brenanii)
Seedling eophylls Pinnate Bifoliolate Pinnate or bipinnate

sometimes do not have spikes arranged in complex synflorescences (e.g. particular
specimens of Ps. bahiana and Ps. contorta). Nonetheless, taken together, the traits
highlighted here provide better recognition of these lineages as distinct genera than
fruit morphology alone.

These results from phylogenetic and morphological analyses provide robust support
for re-circumscription of Pseudopiptadenia as it was traditionally conceived and also
Pityrocarpa. Given that the type species of these two genera are nested in the same clade
and that no morphological traits support the recognition of a narrow circumscription
of Pseudopiptadenia, we subsume the name Pseudopiptadenia under Pityrocarpa, the
oldest validly published generic name (Britton and Rose 1928; Lewis and Lima 1991;
Turland et al. 2018). We assign the remaining Pseudopiptadenia species to the new
genus Marlimorimia.

Key to the genera Parapiptadenia, Pityrocarpa, and Marlimorimia. See Lima et
al. (2022) for a key to all genera of the Stryphnodendron clade

1 Petals reddish (yellowish in Pa. excelsa and Pa. rigida); fruit a legume (dehiscing
along both sutures), the valves plicate above the seeds (except in Pa. excelsa) .
BP UO ete geat trsae Oe org ere ert ones tar eae nee ne sree org ee Parapiptadenia
= Petals white to yellowish or greenish; fruit a follicle (splitting along one suture
only) the valves not plicate: abovest he seed sears. te.d el vsct aeons ee cenerceenenePagnors: 2
2; Petiolar nectary just below or between the first pair of pinnae; spikes solitary

in axils of coevally developing leaves; petals free (united in Pi. leucoxylon) and

248 Leonardo M. Borges et al. / PhytoKeys 205: 239-259 (2022)

glabrous; fruit margins deeply constricted (sinuous in Pi. brenanit)..........04.
sai voinesicbst eth pace ees beak eds ae Ne Ens Roe tT teeta en HeLa Usd ed phat al Taue a Mebetgen Pityrocarpa
- Petiolar nectary between the base and the middle of the petiole; spikes
2-many-fasciculate, the fascicles usually arranged in efoliate terminal pseu-
doracemes or on efoliate nodes below the leaves; petals united and pubescent;
fruit margins straight or shallowly and irregularly sinuous, sometimes con-
SEICLeE WIC SEECStA DOLE. Artis bday lanmehosh ou Annet ars «Raters Marlimorimia

Taxonomy

1. Pityrocarpa (Benth.) Britton & Rose, N. Amer. Fl. 23(3): 190. 1928.

Monoschisma Brenan, Kew Bull. 10(2): 179. 1955, nom. inval., non Monoschisma
Duby, Mém. Soc. Phys. Genéve 19: 294. 1868. Type. Monoschisma leptostachyum
(Benth.) Brenan, syn. nov.

Pseudopiptadenia Rauschert, Taxon 31(3): 559. 1982. Type. Pseudopiptadenia
leptostachya (Benth.) Rauschert, syn. nov.

Basionym. Piptadenia sect. Pityrocarpa Benth., J. Bot. (Hooker) 4: 339. 1842.

Type. Pityrocarpa moniliformis (Benth.) Luckow & R.W. Jobson [= Piptadenia
moniliformis Benth., designated by Britton and Rose 1928].

Description. Unarmed trees or shrubs. Leaves bipinnate; petiole with an
extrafloral nectary between or shortly below the first pair of pinnae; pinnae 1-4
(5) pairs, exceptionally to 10 pairs in Pi. leptostachya; leaflets 1-10 pairs per pinna,
rarely to 20 pairs (Pi. brenanii and Pi. leptostachya), mostly rhomboid sometimes
also asymmetrically elliptical or lanceolate. Inflorescences spikes, solitary in the axils of
coeval leaves, commonly pendulous. Flowers pentamerous; petals free (except possibly
Pi. leucoxylon), glabrous; stamens 10, anther gland present; ovary shortly stipitate
and included within or exserted from the corolla. Fruit a follicle, dehiscing along the
lower suture, flat compressed, mostly moniliform, the margins deeply and regularly
constricted, rarely sinuous margins and shallowly constricted (Pi. brenanii and
occasionally in Pi. leucoxylon); valves stiffly coriaceous. Seeds mostly flat compressed
with a coriaceous testa and a narrow marginal wing, lacking a pleurogram or, less
frequently, ovoid or discoid with a hard, whitish testa, wingless and with a ‘U-
shaped pleurogram (Pi. leucoxylon, Pi. moniliformis and Pi. obliqua); embryo with a
rudimentary plumule (except Pi. brenanii). Seedlings with bifoliolate eophylls.

Distribution. Pityrocarpa is distributed in tropical America, from Mexico to southern
Brazil and Paraguay. Most species occur in the Brazilian Atlantic rainforests (Pi. inaequalis,
Pi. leptostachya, Pi. schumanniana), in the northern Amazonian rainforests (Pi. leucoxylon),
in seasonally dry tropical forests and woodlands in the north-eastern Brazilian Caatinga
(Pi. brenanii, Pi. moniliformis, Pi. obliqua subsp. brasiliensis), western Mexico (Pi. obliqua
subsp. obiqua) or in Venezuelan savannas and Paraguayan Chaco (Pi. moniliformis).

Notes. As circumscribed here, Pityrocarpa includes seven species, all with a mon-
iliform fruit, with the margins deeply constricted between the seeds (Fig. 4). This

Generic re-circumscriptions and a new genus in the Stryphnodendron clade 249
LL
F
E
A ¢ B

G “
Figure 4. Fruits of Pityrocarpa species A Pi. brenanii (from Lewis et al. 1899, NY) B Pi. inaequalis (from
Moreira et al. 3, F) © Pi. leptostachya (from Baez et al. 1174, NY) D Pi. leucoxylon (from de Bruijn 1750,
NY) E Pi. moniliformis (from Nunes 597, HUEEFS) F Pi. obliqua subsp. brasiliensis (from Mori 11837,
NY) G 2%. schumanniana (from Lima 2994, RB).

2cm

D

trait is shared by species formerly included in Pityrocarpa (sensu Jobson and Luckow
2007) and some species previously placed in the genus Pseudopiptadenia (sensu Lewis
and Lima 1991). These two genera had been separated based on seed morphology,
Pityrocarpa characterised by ovoid or discoid seeds with a hard, whitish seed coat and a
‘U-shaped pleurogram, while Pseudopiptadenia included species with flat compressed
and narrowly winged seeds with a coriaceous testa lacking a pleurogram. Pityrocarpa
brenanii and Pi. leucoxylon have fruits with only shallowly sinuous margins, more simi-
lar to species of the genus Marlimorimia.

Besides sharing these fruit traits, Pityrocarpa species also have leaves with few
pinnae (1 to 4 [5] pairs, rarely up to 10 pairs in Pi. leptostachya) and relatively large
thomboid leaflets compared to species of Marlimorimia. One exception are the leaves of
Pi. brenanii, which are similar to those of M. bahiana. All species of Pityrocarpa present
an extrafloral nectary between or shortly below the first pair of pinnae, in contrast to
species of Marlimorimia that have the nectary below mid-petiole, frequently close to
the pulvinus.

250 Leonardo M. Borges et al. / PhytoKeys 205: 239-259 (2022)

Floral traits, although previously disregarded as being generically diagnostic
in the group, provide further evidence for the distinction between Pityrocarpa and
Marlimorimia. The solitary inflorescence spikes in the axils of coevally developing
leaves in Pityrocarpa contrast with the more complex synflorescences of Marlimorimia
(Fig. 3; see notes under Marlimorimia). All species of Pityrocarpa have free and glabrous
petals, except for Pi. leucoxylon, in which the petals are connate for a little over 1 mm
(Barneby and Grimes 1984).

Lima (1985) and Lewis and Lima (1991) provided additional information on
embryos and seedlings that are potentially useful for distinguishing Pityrocarpa from
Marlimorimia. Embryos of Pityrocarpa species have a rudimentary plumule, while
in Marlimorimia, the plumule is developed and multifid. This seems to be correlated
with seedling morphology as the studied species of Pityrocarpa have bifoliolate eo-
phylls and those of Marlimorimia species have pinnate or bipinnate eophylls (Lewis
and Lima 1991). Pityrocarpa brenanii, however, has embryo morphology more simi-
lar to that reported for species of Marlimorimia (Lewis and Lima 1991).

1.1. Pityrocarpa brenanii (G.P. Lewis & M.P. Lima) L.P. Queiroz & L.M. Borges,
comb. nov.
urn:lsid:ipni.org:names:77303780-1

Basionym. Pseudopiptadenia brenanii G.P. Lewis & M.P. Lima, Arch. Jard. Bot. Rio de
Janeiro 30: 50-51. 1991.

Type. Brasil, Bahia, Harley et al. 21346 (holotype CEPEC; isotypes BR, K, M,
MBM, MEXU, NY, RB, US).

1.2. Pityrocarpa inaequalis (Benth.) L.P. Queiroz & Marc.F. Simon, comb. nov.
urn:lsid:ipni.org:names:7730378 1-1

Monoschisma inaequale (Benth.) Brenan, Kew Bull. 10(2): 179. 1955.
Pseudopiptadenia inaequalis (Benth.) Rauschert, Taxon 31(3): 559. 1982.

Basionym. Piptadenia inaequalis Benth., J. Bot. (Hooker) 4: 339. 1842.
Type. Brazil, Rio de Janeiro, Pohl 1386 (lectotype K 000504704, designated here;
isolectotype K 000504706).

1.3. Pityrocarpa leptostachya (Benth.) L.P. Queiroz & P.G. Ribeiro, comb. nov.
urn:lsid:ipni.org:names:77303782-1

Monoschisma leptostachyum (Benth.) Brenan, Kew Bull. 10(2): 179. 1955.
Pseudopiptadenia leptostachya (Benth.) Rauschert, Taxon 31(3): 559. 1982.

Generic re-circumscriptions and a new genus in the Stryphnodendron clade 251

Basionym. Piptadenia leptostachya Benth., J. Bot. (Hooker) 4: 339. 1842.

Type. Brasil, Sellow s.n. (Lectotype K 000504709, designated here; isolectoypes F
0360957F [fragment], K 000504710, TUB 009699).

Note. Lewis and Lima (1991) unintentionally lectotypified this name by indicat-
ing the holotype to be at B and the isotype to be at K. However, the B specimen was
destroyed and, hence, cannot serve as a lectotype. Moreover, K holds two duplicates
of an un-numbered Sellow collection. Here, we chose the one previously belonging to
Bentham’s herbarium as the lectotype.

1.4. Pityrocarpa leucoxylon (Barneby & J.W. Grimes) Luckow & R.W. Jobson,
Syst. Bot. 32(3): 573. 2007.

Basionym. Piptadenia leucoxylon Barneby & J.W. Grimes, Brittonia 36(3): 236-238.
1984.
Type. Venezuela: Bolivar, de Bruijn 1750 (holotype NY; isotypes MO, VEN, US).

1.5. Pityrocarpa moniliformis (Benth.) Luckow & R.W. Jobson, Syst. Bot. 32(3):
573. 2007.

Stryphnodendron piptadenioides ELM.O. Martins, Leandra 5(6): 90. 1975. Type. Brazil.
Pernambuco, “lectum in silva pluviali ad S. José Belmonte”, Mata da Mina, 29 Oct
1971, Ramatlho 52 (holotype RFA 17173) .

Stryphnodendron consimile E.M.O. Martins, Leandra 5(6): 92. 1975. Type. Brazil. Pi-
aui, “habitat in caatinga ad Paulistana’, Fazenda Altamira, 04 Nov 1974, Lima
1330 (holotype RFA 17172).

Basionym. Piptadenia moniliformis Benth., J. Bot. (Hooker) 4: 339. 1842.
Type. Brazil, Bahia, Serra de Jacobina, Blanchet 2701 (lectotype K 000090193,
designated here; isolectotypes F, K 000205897, MO, NY 00003233).

1.6. Pityrocarpa obliqua (Pers.) Brenan, Kew Bull. 10(2): 176. 1955.

Acacia thibaudiana DC., Prodr. 2: 456. 1825.
Piptadenia obliqua (Pers.) J.F. Macbr., Contr. Gray Herb. 59: 17. 1919.

Basionym. Sophora obliqua Pers., Syn. P/. 1: 452. 1805.
Type. “Amer. australi?”, Herb. D. Thibaud. (not located).

1.6.1. Pityrocarpa obliqua subsp. obliqua

252 Leonardo M. Borges et al. / PhytoKeys 205: 239-259 (2022)

1.6.2. Pityrocarpa obliqua subsp. brasiliensis (G.P. Lewis) Luckow & R.W. Job-
son, Syst. Bot. 32(3): 573. 2007.

Basionym. Piptadenia obliqua subsp. brasiliensis G.P. Lewis, Kew Bull. 46(1): 160-
162. 1991.
Type. Brazil, Bahia, Mori et al. 9519 (holotype CEPEC; isotypes HUEFS, K, NY).

1.7. Pityrocarpa schumanniana (Taub.) L.P. Queiroz & L.M. Borges, comb. nov.
urn:lsid:ipni.org:names:77303784- 1

Pseudopiptadenia schumanniana (Taub.) G.P. Lewis & M.P. Lima, Arch. Jard. Bot. Rio
de Janeiro 30: 53. 1991.

Basionym. Piptadenia schumanniana Yaub., Flora 75: 75. 1892.

Type. Brazil, “Brasilia austro-orientale”, Rio de Janeiro, Glaziou 13774 (lectotype
R 00008369, designated here; isolectotypes A 00064056, F 0058675, K 000504703,
MPU 016109, NY 00003244, NY 00003245, US 00001018, US 00997081).

2. Marlimorimia L.P. Queiroz, L.M. Borges, Marc.F. Simon & PG. Ribeiro, gen. nov.
urn:lsid:ipni.org:names:77303785-1

Newtonia sect. Neonewtonia Burkart, Fl. Il. Catarin. fasc. LEGU: 285. 1979, syn. nov.
Type. Newtonia nitida (Benth.) Brenan (= Marlimorimia contorta (DC.) L.P. Quei-
roz & P.G. Ribeiro).

Diagnosis. Marlimorimia shares with Pityrocarpa the follicle, a fruit dehiscing along the
lower suture only, and flat, compressed winged seeds, which lack a pleurogram. It can be
differentiated from Pityrocarpa by the position of the extrafloral nectary on the petiole
(from the base to the mid-petiole in Marlimorimia vs. between or just below the first pair
of pinnae in Pityrocarpa); inflorescence spikes clustered in terminal pseudoracemes or in
fascicles at efoliate nodes, surpassed by mature leaves (vs. solitary spikes in the axils of coeval
leaves); petals united and joined into a gamopetalous corolla (vs. petals free and glabrous);
and fruits with margins straight to shallowly sinuous (vs. margins deeply constricted).

Type. Marlimorimia contorta (DC.) L.P. Queiroz & P.G. Ribeiro

Description. Unarmed trees. Leaves bipinnate; petiole with an extrafloral nectary
well below the first pair of pinnae, close to the pulvinus, always below mid-petiole;
pinnae 5—10 to many pairs per leaf (2—3 pairs in VM. colombiana and 3-5 in M. bahiana);
leaflets mostly > 10 pairs per pinna, (6-8 in M. colombiana), mostly oblong to linear
from an asymmetrical base, rarely rhomboid (M. bahiana). Inflorescences spikes, grouped
in fascicles, these being arranged in terminal pseudoracemes or forming clusters below
the coeval leaves. Flowers pentamerous; petals united into a gamopetalous corolla,
pubescent; stamens 10, anther gland present; ovary shortly stipitate and included

Generic re-circumscriptions and a new genus in the Stryphnodendron clade 253

or exserted from the corolla. Fruit a follicle, dehiscing along the lower suture, flat
compressed, straight, curved or longitudinally twisted, the margins usually straight,
rarely irregularly sinuous and only becoming constricted where the seeds fail to
develop (M. bahiana and M. warmingii), valves coriaceous, thin or thick. Seeds flat
compressed with a coriaceous testa, presenting a narrow or somewhat wider marginal
wing, pleurogram lacking; embryo with a developed, multifid plumule (unknown in
M. colombiana and M. pittieri). Seedlings with pinnate or bipinnate eophylls (unknown
in M. bahiana, M. colombiana and M. pittieri).

Distribution. Marlimorimia comprises six species with a bicentric distribution in
the two main areas of tropical humid forests in South America. Three species occur
in eastern Brazil, two of which are restricted to the Atlantic wet forests (Marlimorima
bahiana and M. warmingii) and M. contorta, which extends to inland semi-
deciduous forests. The three other species are distributed in northern South America.
Marlimorimia psilostachya is widely distributed across Amazonia, sparsely extending to
Central America (Costa Rica) and M. colombiana and M. pittieri have restricted ranges
in Colombia and Venezuela, respectively.

Etymology. The genus Marlimorimia is named in honour of Dr. Marli Pires Morim,
taxonomist at the Rio de Janeiro Botanical Garden, for her outstanding contribution to
our knowledge of the diversity and taxonomy of Brazilian mimosoid legumes.

Notes. The new genus Marlimorimia is proposed to accommodate a monophyletic
group of species, previously classified in Pseudopiptadenia (sensu Lewis and Lima 1991;
Luckow 2005), but which could not retain the genus name, because its type species is
now included in Pityrocarpa.

Besides the molecular phylogenetic evidence, morphology also supports recognition
of Marlimorimia as distinct from Pityrocarpa. Marlimorimia brings together most of the
species formerly placed in Pseudopiptadenia which have multipinnate leaves, small oblong
to linear leaflets and fruits with straight (or shallowly sinuous) margins. Marlimorimia
bahiana and M. colombiana, however, have leaves with few pinnae and rhomboid leaflets.

Species of Marlimorimia have more complex inflorescences than those of Pityrocar-
pa. While the spikes of Pityrocarpa are solitary in the axils of coevally developing leaves,
Marlimorimia species have spikes in fascicles of 2-3, which are arranged in terminal
efoliate pseudoracemes or clustered on nodes below mature leaves (Fig. 3). Sometimes,
as leaves expand, Marlimorimia synflorescences may resemble those of Pityrocarpa and
Parapiptadenia (e.g. particular specimens of M. contorta such as Hatschbach 50149
[NY]). Nonetheless, flowers of Marlimorimia have pubescent petals united into a
gamopetalous corolla (vs. free glabrous petals in the majority of Pityrocarpa species).

Two types of fruits are found in Marlimorimia (Fig. 5). Some species have
long linear fruits, frequently curved or longitudinally twisted with straight margins
(M. colombiana, M. contorta, M. pittieri and M. psilostachya), while M. bahiana and
M. warmingii have oblong fruits with shallowly sinuous margins. The valves of the
fruits are woody, although usually thin, becoming thicker and harder in M. warmingii.

The seeds of Marlimorimia, although superficially similar to those of most species
of Pityrocarpa, have embryos with multifid plumules that result in seedlings with pin-
nate or bipinnate eophylls (Lima 1985; Lewis and Lima 1991).

254 Leonardo M. Borges et al. / PhytoKeys 205: 239-259 (2022)

B —<
A C 2cm E

Figure 5. Fruits of Marlimorimia species A Marlimorimia bahiana (from Amorim 1009, NY)
B Marlimorimia colombiana (from Killip 16268, NY) © Marlimorimia contorta (from Gomes 257, NY)
D Marlimorimia pittieri (from Guevara 1264, F) E Marlimorimia psilostachya (from Rabelo 2753, NY)
F Marlimorimia warmingii (from Nunes et al. 2, NY).

2.1. Marlimorimia bahiana (G.P. Lewis & M.P. Lima) L.P. Queiroz & L.M. Borg-
es, comb. nov.

urn:lsid:ipni.org:names:77303786-1

Basionym. Pseudopiptadenia bahiana G.P. Lewis & M.P. Lima, Arch. Jard. Bot. Rio de
Janeiro 30: 54-55. 1991.

Type. Brasil, Bahia, Mori & King 12223 (holotype CEPEC; isotypes HUEFS, K,
NY, RB).

Generic re-circumscriptions and a new genus in the Stryphnodendron clade 250

2.2. Marlimorimia colombiana (Britton & Killip) L.P. Queiroz & Marc.F. Simon,
comb. nov.

urn:lsid:ipni.org:names:77303787-1
Pseudopiptadenia colombiana (Britton & Killip) G.P. Lewis

Basionym. Stryphnodendron colombianum Britton & Killip, Ann. New York Acad. Sci.
35 (3)al 5a. 1936.

Type. Colombia, Santander, Killip & Smith 16268 (holotype NY 00003356; iso-
types A, GH, US).

Notes. In the absence of phylogenetic evidence, the petiolar extrafloral nectaries
located at mid-petiole and fruits with straight to shallowly sinuous margins support
the transfer of Pseudopiptadenia colombiana to Marlimorimia.

2.3. Marlimorimia contorta (DC.) L.P. Queiroz & P.G. Ribeiro, comb. nov.
urn:lsid:ipni.org:names:77303788- 1

Piptadenia nitida Benth., J. Bot. (Hooker) 4: 336. 1842.

Piptadenia contorta (DC.) Benth., Trans. Linn. Soc. Lond. 30: 368. 1875.

Newtonia nitida (Benth.) Brenan, Kew. Bull. 10 (2): 182. 1955.

Newtonia contorta (DC.) Burkart, FI. Il. Catarin. fasc. LEGU: 289. 1979.

Pseudopiptadenia contorta (DC.) G.P. Lewis & M.P. Lima, Arch. Jard. Bot. Rio de
Janeiro 30: 57. 1991.

Basionym. Acacia contorta DC., Prodr. 2: 470. 1825.
Type. Brasil, Rio de Janeiro, Raddi s.n. (lectotype FI, designated here).

2.4. Marlimorimia pittieri (Harms) L.P. Queiroz & L.M. Borges, comb. nov.
urn:lsid:ipni.org:names:77303789- 1

Piptadenia similis Britton & Killip, Ann. New York Acad. Sci. 35(3): 156. 1936.
Holotype Colombia, Barranquilla, Elias 263 (US).
Pseudopiptadenia pittieri (Harms) G.P. Lewis, Kew Bull. 46(1): 118. 1991.

Basionym. Piptadenia pittieri Harms, Notizbl. Bot. Gart. Berlin-Dahlem 8(71): 51-
BAe be PA ls

Type. Venezuela, Carabobo, Pittier 8859 (lectotype US 00001013, designated
here; isolectotypes GH 00064052, NY 00003236).

Notes. Although Pseudopiptadenia pittieri was not included in the phylogenetic
analyses, the presence of extrafloral nectaries at the base of the petiole, spikes arranged
in pseudoracemes and fruits with straight margins support its transfer to Marlimorimia.

256 Leonardo M. Borges et al. / PhytoKeys 205: 239-259 (2022)

2.5. Marlimorimia psilostachya (DC.) L.P. Queiroz & Marc.F. Simon, comb. nov.
urn:lsid:ipni.org:names:77303790-1

Piptadenia psilostachya (DC.) Benth., J. Bot. (Hooker) 4: 336. 1842.

Piptadenia suaveolens Migq., Linnaea 18: 589-590. 1845. Type Surinam, Bergendaal,
Focke 936 (holotype U).

Newtonia psilostachya (DC.) Brenan, Kew. Bull. 10 (2): 182. 1955.

Newtonia suaveolens (Miq.) Brenan, Kew. Bull. 10 (2): 182. 1955.

Pseudopiptadenia psilostachya (DC.) G.P. Lewis & M.P. Lima, Arch. Jard. Bot. Rio de
Janeiro 30: 55. 1991.

Pseudopiptadenia suaveolens (Migq.) J.W. Grimes, Brittonia 45(1): 27. 1993.

Basionym. Acacia psilostachya DC., Prodr. 2: 457. 1825.

Type. French Guiana, Cayenne, Martin 2 (lectotype K 000504699, designated by
Lewis & Lima 1991; isolectotype P 02930999).

Notes. Contrary to Grimes (1993), who recognised Pseudopiptadenia psilostachya
and Ps. suaveolens as distinct species, we agree with Lewis and Lima (1991) on the syn-
onymisation of Ps. suaveolens under M. psilostachya. These plants grow sympatrically and
the traits used by Grimes (1993) to support recognition of two species are too variable
to be diagnostic.

2.6. Marlimorimia warmingii (Benth.) L.P. Queiroz & P.G. Ribeiro, comb. nov.
urn:lsid:ipni.org:names:7730379 1-1

Piptadenia glaziovii Harms, Repert. Spec. Nov. Regni Veg. 17: 203. 1921. Type. Brasil,
Rio de Janeiro, Serra da Estrela, Glaziou 8440 (lectotype K, designated by Lewis
and Lima 1991).

Newtonia glaziovii (Harms) Burkart ex Barth & Yoneshigue, Mem. Inst. Oswaldo Cruz
64: 102. 1966.

Newtonia warmingii (Benth.) G.P. Lewis, Legumes of Bahia p. 111. 1987.

Pseudopiptadenia warmingii (Benth.) G.P. Lewis & M.P. Lima, Arch. Jard. Bot. Rio de
Janeiro 30: 54. 1991.

Basionym. Mimosa warmingii Benth., Trans. Linn. Soc. London 30(3): 413. 1875.
Type. Brasil, Minas Gerais, Lagoa Santa, Warming s.n. (lectotype K 000504702,
designated by Lewis and Lima 1991).

Acknowledgements

We thank fellow plant collectors, particularly Haroldo de Lima, Danilo Neves and Do-
mingos Cardoso, for providing silica-dried leaf samples used in our phylogenetic analy-
sis. We also thank Marli Morim for her valuable inputs to the manuscript, Thais Cury de

Generic re-circumscriptions and a new genus in the Stryphnodendron clade 257

Barros for a discussion on anther gland morphology and Gwilym Lewis, an anonymous
reviewer and particularly Colin Hughes for their suggestions. L.P. Queiroz acknowledg-
es support from CNPq (processes 303585/2016-1 and 440487/2015-3) and FAPESB
(processes APP 096/2016 and PTX0004/2016). P.G. Ribeiro acknowledges a Ph.D.
grant from Coordenacao de Aperfeigoamento de Pessoal de Nivel Superior (CAPES;
process 5414130) and the GM and REFLORA project grants from CNPq (processes
130515/2010-8 and 563533/2010-2). M.F. Simon acknowledges support from CNPq
(305570/2021-8). The use of DNA from the Brazilian species is authorised by SISGEN.

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Supplementary material |

Voucher information and GenBank accession numbers (internal transcribed spac-

et-ITS sequences) for taxa used in this study

Authors: Leonardo M. Borges, Peter W. Inglis, Marcelo F Simon, Pétala Gomes

Ribeiro, Luciano P. de Queiroz

Data type: excel file

Explanation note: Voucher information and GenBank accession numbers for taxa used
in this study.

Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODDbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Link: https://doi.org/10.3897/phytokeys.205.82275.suppl1